Length-Tension Relationship Length-Tension Relationship • Amount of tension and force of contraction depends on how stretched or contracted muscle was before it’s stimulated Optimum resting length z z • If overly contracted at rest, a weak contraction results Overly contracted z • if too stretched before stimulated, a weak contraction results z Overly stretched z – If Sarcomre is < 60% or >175% of original length develop no tension z 1.0 • optimum resting length produces greatest force when muscle contracts Tension generated 0.5 – CNS adjusts our muscle tone – state of partial contraction – maintains optimum length – i.e., always ready for action 0.0 1.0 2.0 3.0 Sarcomere length (µm) before stimulation 4.0 11-1 11-2 Behavior of Whole Muscles Phases of twitch contraction • Latent Period: time required for excitation, excite/coupling and tension - internal tension, no shortening • twitch – a quick cycle of contraction and relaxation • Contraction phase – sliding filamants - external tension • subthreshold electrical stimulus causes = no contraction • Relaxation phase - SR reabsorbs Ca+2, myosin releases actin - - tension declines – muscle returns to resting length – twitch lasts from 7 to 100 msec Latent period Time of stimulation Time Figure 11.13 11-3 at threshold intensity and above - a twitch is produced – twitches caused by increased voltage are no stronger than those at threshold – i.e. muscles follow all-or-none law (contracting to its maximum or not at all) Stimulus voltage subthreshold stimulus – no contraction at all • 11-4 Recruitment and Stimulus Intensity Contraction Strength of Twitches • Relaxation phase Contraction phase Muscle tension • threshold - minimum voltage necessary to generate an action potential in the muscle fiber and produce a contraction Threshold 1 2 3 4 5 6 7 8 9 Stimuli to nerve Figure 11.14 Proportion of nerve fibers excited not exactly true – twitches vary in strength depending upon: • stimulus frequency - stimuli arriving closer together produce stronger twitches • concentration of Ca+2 in sarcoplasm can vary the frequency • how stretched muscle was before it was stimulated • temperature : warmed-up muscle contracts more strongly – enzymes work more quickly • pH: if low in sarcoplasm weakens contraction - fatigue • hydration of muscle affects overlap of actin & myosin Maximum contraction Tension • 1 2 3 4 5 6 7 8 9 Responses of muscle • stimulating nerve with higher and higher voltages produces stronger contractions – higher voltages excite more nerve fibers in the motor nerve which stimulates more motor units to contract 11-5 • recruitment or multiple motor units (MMU) summation – bringing more motor units into play 11-6 1 Twitch Strength & Stimulus Frequency Treppe: stimulus right after relaxation following tension is greater than previous Twitch Muscle twitches Muscle develops tension but does not shorten Muscle shortens, tension remains constant Muscle lengthens while maintaining tension Movement Movement No movement Stimuli (a) (b) Incomplete tetanus: stimuli before muscle relaxes = summation of twitches – rapid cycles of contr. & relax. Tension rises! Complete tetanus: freq so great there is no relaxation (a) Isometric contraction (b) Isotonic concentric contraction (c) Isotonic eccentric contraction Fatigue 11-7 (c) (d) Isometric and Isotonic Phases of Contraction Muscle Metabolism • all muscle contraction depends on ATP Length or Tension • ATP supply depends on availability of: Muscle tension – oxygen – organic energy sources such as glucose and fatty acids Muscle length • two main pathways of ATP synthesis Isometric phase aerobic respiration • produces far more ATP • less toxic end products (CO2 and water) • requires a continual supply of oxygen – anaerobic fermentation • produce ATP in the absence of O2 • yields little ATP and lactic acid, a major factor in muscle fatigue Isotonic phase Time • at the beginning of contraction – isometric phase – muscle tension rises but muscle does not shorten • when tension overcomes resistance of the load – tension levels off • muscle begins to shorten and move the load – isotonic phase Overview of ATP Production Glycolysis Stages of glucose oxidation 11-9 11-10 Modes of ATP Synthesis During Exercise Glucose 2 ADP + 2 Pi 2 ATP Pyruvic acid Anaerobic fermentation No oxygen available Lactic acid Aerobic respiration Oxygen available Glycogen: - Stored in Liver, Skeletal muscles, heart - Can be changed into glucose CO2 + H2O Mitochondrion 36 ADP + 36 Pi 0 10 seconds 40 seconds Duration of exercise Repayment of oxygen debt Mode of ATP synthesis At rest most ATP generated by aerobic respir. of fatty acids - During short intense exercise Aerobic respiration using oxygen from Myoglobin - ATP is used up quickly until respiratory system catches up – until then Phosphcreatine!!!!! – Until it gets used up!!!! Now muscles using Glycogen– lactic acid system (anaerobic fermentation) Aerobic respiration supported by cardiopulmonary Bringing O2 to muscles 36 ATP • ATP consumed within 60 seconds of formation 2 Immediate Energy Needs Fatigue ADP ADP • Causes of muscle fatigue – ATP synthesis declines as glycogen is consumed – ATP shortage slows down the Na+ - K+ pumps – lactic acid lowers pH of sarcoplasm • inhibits enzymes involved in contraction, • Inhibits enzymes used in ATP synthesis – release of K+ with each action potential causes the accumulation of extracellular K+ • hyperpolarizes the cell and makes the muscle fiber less excitable – motor nerve fibers use up their Achetelcholine – CNS fatigues ??????? less signal output to skeletal muscles Pi Myokinase ATP AMP ADP Creatine phosphate Pi Figure 11.19 Creatine Creatine kinase ATP 11-13 11-14 Oxygen Debt Endurance • endurance – ability to maintain high-intensity exercise for > 4 to 5 minutes – determined in large part by one’s maximum oxygen uptake (VO2max) – maximum oxygen uptake – point at which rate of oxygen consumption reaches a plateau and does not increase further with added workload • When exercise stops, rate of oxygen uptake does not immediately return to pre-exercise levels • Because oxygen debt accumulated during exercise: – When oxygen is withdrawn from hemoglobin and myoglobin – And because of O2 needed for metabolism of lactic acid produced by anaerobic respiration - Turn it back to pyruvic acid then to glycogen 11-15 12-49 Physiological Classes of Muscle Fibers FG and SO Muscle Fibers Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. • slow oxidative (SO), slow-twitch, red, or type I fibers – Lots of mitochondria, myoglobin and capillaries - deep red color – adapted for aerobic respiration - - slow to fatigue – long lasting twitch (~100 msec) • fast glycolytic (FG), fast-twitch, white, or type II fibers – few mitochondria, myoglobin, and blood capillaries - pale appearance – adapted for quick responses - - fast to fatigue (lactic acid) – rich in enzymes of phosphagen and glycogen-lactic acid systems generate lactic acid causing fatigue – fast twitches (~ 7.5 msec) FG SO • ratio of different fiber types have genetic predisposition 11-17 Figure 11.20 11-18 3 Strength and Conditioning Strength and Conditioning • muscles can generate more tension than bones and tendons can withstand • resistance training (weight lifting) – – – – • muscular strength depends on: – primarily on muscle size – fascicle arrangement contraction of a muscles against a load that resist movement Couple times a week Hypertrophy more myofilaments and myofibrils synthesized • pennate are stronger than parallel, and parallel stronger than circular • endurance training (aerobic exercise) – size of motor units – multiple motor unit summation – recruitment of many motor units – temporal summation • nerve impulses usually arrive at a muscle in a series of closely spaced action potentials – length – tension relationship (resting muscle at optimal length) – improves fatigue resistant muscles – slow twitch fibers produce more mitochondria, glycogen, and acquire a greater density of blood capillaries – improves skeletal strength – increases red blood cell count and oxygen transport capacity of the blood – enhances cardiovascular, respiratory, and nervous systems – fatigue 11-19 11-20 Cardiac Muscle Cardiac Muscle • characteristics of cardiac muscle cells: • Location? • Function? – striated but myocytes (cardiocytes) shorter and thicker than skelatal • Required properties of cardiac muscle – intercalated discs • electrical gap junctions allow each cell to directly stimulate its neighbors • mechanical junctions that keep the cells from pulling apart – Autorhythmic cells – contraction with regular rhythm Ca+ induced Ca+ release from Sarcoplasmic reticulum – muscle cells of each chamber must contract in unison Damaged cardiac muscle cells repair by fibrosis – contractions must last long enough to expel blood • a little mitosis observed following heart attacks • not in significant amounts to regenerate functional muscle – must be highly resistant to fatigue 11-21 Cardiac Muscle 11-22 Smooth Muscle • myocytes have a fusiform shape & one nucleus • no visible striations • Actin and myosin do not overlap • z discs absent • No troponin • Cytoplasm has: – autorhythmic cells!!!! – autonomic nervous system can influence it – slow twitches!! • maintains tension for about 200 to 250 msec – uses aerobic respiration almost exclusively • rich in myoglobin and glycogen • has especially large mitochondria - dense bodies = Like z disks – protein plaques on the inner face of the plasma – cytoskeleton of intermediate filaments: attach to the membrane plaques and dense bodies – highly fatigue resistant 11-23 • Ca2+ needed for muscle contraction comes from ECF • ANS controlled • capable of mitosis (hyperplasia) Plaque Intermediate filaments of cytoskeleton Actin filaments Dense body Myosin 11-24 4 2 Types of Smooth Muscle 2 Types of Smooth Muscle • single-unit smooth muscle (visceral muscle) Autonomic nerve fibers • multiunit smooth muscle Autonomic nerve fibers – myocytes are electrically coupled to each other by gap junctions – occurs in some of the largest arteries and pulmonary air passages, in piloerector muscles of hair follicle, and in the iris of the eye – stimulate each other and a large number of cells contract as a single unit – i.e., each cell does NOT have to be stimulated directly by a motor neuron Synapses – Single neuron stimulates many myocytes • i.e., like a motor unit Varicosities Gap junctions (a) Multiunit smooth muscle Figure 11.21a (b) Single-unit smooth muscle 11-25 Figure 11.21b 11-26 Stimulation of Smooth Muscle Layers of Visceral Smooth Muscle • Involuntary and can contract without nervous stimulation!! – chemical stimuli: hormones, carbon dioxide, low pH, and oxygen deficiency – in response to stretch – Cold stimulus – (aeola and scrotum) – Stretching - peristalsis • most innervated by ANS nerve fibers – can trigger and modify contractions – Neurotransmitters: acetylcholine or norepinephrine Mucosa: Epithelium Lamina propria Muscularis mucosae Muscularis externa: Circular layer Longitudinal layer Figure 11.22 11-27 Stimulation of Smooth Muscle 11-28 Contraction and Relaxation • contraction is triggered by Ca+2, ATP, and sliding thin past thick myofilaments Autonomic nerve fiber • contraction begins in response to Ca+2 that enters the cell from ECF – voltage, ligand, and mechanically-gated (stretching) Ca+2 channels open (Ca+2 enters cell) Varicosities • Ca+ binds to calmodulin – activates myosin light-chain kinase – adds phosphate to myosin head – This activates myosin ATPase - hydrolyzing ATP – allowing myosin head to move Actin – thick filaments pull on thin ones, this pulls on dense bodies and membrane plaques Mitochondrion Synaptic vesicle Single-unit smooth muscle Figure 11.23 11-29 11-30 5 Please note that due to differing operating systems, some animations will not appear until the presentation is viewed in Presentation Mode (Slide Show view). You may see blank slides in the “Normal” or “Slide Sorter” views. All animations will appear after viewing in Presentation Mode and playing each animation. Most animations will require the latest version of the Flash Player, which is available at http://get.adobe.com/flashplayer. enzyme Component of myosin 12-81 Stretching Smooth Muscle Plaque • stretch can open mechanically-gated calcium channels in the sarcolemma causing contraction – peristalsis – waves of contraction brought about by food distending the esophagus or feces distending the colon • propels contents along the organ Intermediate filaments of cytoskeleton • stress-relaxation response (receptive relaxation) -helps hollow organs gradually fill (urinary bladder) – when stretched, tissue briefly contracts then relaxes – helps prevent emptying while filling Actin filaments Dense body Myosin (b) Contracted smooth muscle cells Figure 11.24 (a) Relaxed smooth muscle cells 11-33 11-34 Contraction and Stretching • smooth muscle contracts forcefully even when greatly stretched – Remember what happens when Skeletal muscle is stretched? • plasticity – the ability to adjust its tension to the degree of stretch – a hollow organ such as the bladder can be greatly stretched yet not become flabby when it is empty 11-35 6